Significance of summer fog and overcast for drought stress and ecological functioning of coastal California endemic plant species

Aim Fog drip is a crucial water source for plants in many ecosystems, including a number of global biodiversity hotspots. In California, dozens of rare, drought‐sensitive plant species are endemic to coastal areas where the dominant summer moisture source is fog. Low clouds that provide water to these semi‐arid ecosystems through fog drip can also sharply reduce evaporative water losses by providing shade. We quantified the relative hydrological importance of cloud shading vs. fog drip. We then examined how both factors influence the range dynamics of an apparently fog‐dependent plant species spanning a small‐scale cloud gradient. Location The study area is on Santa Cruz Island off the coast of southern California. It is near the southern range limit of bishop pine (Pinus muricata D. Don), a tree endemic to the coasts of California and Baja, Mexico. Methods We measured climate across a pine stand along a 7 km, coastal–inland elevation transect. Short‐term (1–5 years) monitoring and remote sensing data revealed strong climatic gradients driven primarily by cloud cover. Long‐term (102 years) effects of these gradients were estimated using a water balance model. Results We found that shade from persistent low clouds near the coast reduced annual drought stress by 22–40% compared with clearer conditions further inland. Fog drip at higher elevations provided sufficient extra water to reduce annual drought stress by 20–36%. Sites located at both high elevation and nearer the coast were subject to both effects. Together, these effects reduced average annual drought stress by 56% and dramatically reduced the frequency of severe drought over the last century. At lower elevation (without appreciable fog drip) and also near the inland edge of the stand (with less cloud shading) severe droughts episodically kill most pine recruits, thereby limiting the local range of this species. Main conclusions Persistent cloud shading can influence hydrology as much as fog drip in cloud‐affected ecosystems. Understanding the patterns of both cloud shading and fog drip and their respective impacts on ecosystem water budgets is necessary to fully understand past species range shifts and to anticipate future climate change‐induced range shifts in fog‐dependent ecosystems.     

The influence of summertime fog and overcast clouds on the growth of a coastal Californian pine: a tree-ring study

The coast of California is home to numerous rare, endemic conifers and other plants that are limited in distribution by drought sensitivity and the summer-dry climate that prevails across most of the state. Ecologists have long assumed that some coastal plant populations survived the early Pleistocene transition to a warmer and drier environment because they benefit from frequent fog and stratus clouds that provide water and shade during the rainless summer. One such population is that of Torrey pine (Pinus torreyana ssp. Insularis) on Santa Rosa Island in Channel Islands National Park. Here we report that the tree-ring width record from this population indicates strong growth sensitivities to summer fog drip and cloud shading. We quantified the effects of summer cloud cover by comparing ring-width indices to coastal airport cloud-frequency records (1944–2004). For the first time observed, summertime cloud frequency correlated positively with ring-width indices, regardless of whether the effect of rainfall was first removed from the ring-width record. The effect of ground-level fog was strongest in July early mornings (03:00 PST, R ² = 0.262, P < 0.0002). The effect of clouds high enough to provide shade but not fog water was also strongest in July, but climbed steadily throughout the day before becoming strongest in late afternoon (16:00–18:00 PST, R ² = 0.148, P < 0.004). Correlations were substantially stronger in years with higher soil moisture, suggesting that growth response to summer clouds is strongly affected by pre-summer rainfall. A change in the height and/or timing of coastal cloud formation with climate change would likely affect this and other populations of California’s coastal vegetation.     

Fog in the California redwood forest: ecosystem inputs and use by plants

Fog has been viewed as an important source of moisture in many coastal ecosystems, yet its importance for the plants which inhabit these ecosystems is virtually unknown. Here, I report the results of a 3-year investigation of fog inputs and the use of fog water by plants inhabiting the heavily fog inundated coastal redwood (Sequoia sempervirens) forests of northern California. During the study period, 34%, on average, of the annual hydrologic input was from fog drip off the redwood trees themselves (interception input). When trees were absent, the average annual input from fog was only 17%, demonstrating that the trees significantly influence the magnitude of fog water input to the ecosystem. Stable hydrogen and oxygen isotope analyses of water from fog, rain, soil water, and xylem water extracted from the dominant plant species were used to characterize the water sources used by the plants. An isotopic mixing model was employed to then quantify how much fog water each plant used each month during the 3-year study. In summer, when fog was most frequent, ～19% of the water within S.sempervirens, and ～66% of the water within the understory plants came from fog after it had dripped from tree foliage into the soil; for S.sempervirens, this fog water input comprised 13–45% of its annual transpiration. For all plants, there was a significant reliance on fog as a water source, especially in summer when rainfall was absent. Dependence on fog as a moisture source was highest in the year when rainfall was lowest but fog inputs normal. Interestingly, during the mild El Niño year of 1993, when the ratio of rainfall to fog water input was significantly higher and fog inputs were lower, both the proportion and coefficient of variation in how much fog water was used by plants increased. An explanation for this is that while fog inputs were lower than normal in this El Niño year, they came at a time when plant demand for water was highest (summer). Therefore, proportional use of fog water by plants increased. The results presented suggest that fog, as a meteorological factor, plays an important role in the water relations of the plants and in the hydrology of the forest. These results demonstrate the importance of understanding the impacts of climatic factors and their oscillations on the biota. The results have important implications for ecologists, hydrologists, and forest managers interested in fog-inundated ecosystems and the plants which inhabit them.     

Seasonal and episodic moisture controls on plant and microbial contributions to soil respiration

Moisture inputs drive soil respiration (SR) dynamics in semi-arid and arid ecosystems. However, determining the contributions of root and microbial respiration to SR, and their separate temporal responses to periodic drought and water pulses, remains poorly understood. This study was conducted in a pine forest ecosystem with a Mediterranean-type climate that receives seasonally varying precipitation inputs from both rainfall (in the winter) and fog-drip (primarily in the summer). We used automated SR measurements, radiocarbon SR source partitioning, and a water addition experiment to understand how SR, and its separate root and microbial sources, respond to seasonal and episodic changes in moisture. Seasonal changes in SR were driven by surface soil water content and large changes in root respiration contributions. Superimposed on these seasonal patterns were episodic pulses of precipitation that determined the short-term SR patterns. Warm season precipitation pulses derived from fog-drip, and rainfall following extended dry periods, stimulated the largest SR responses. Microbial respiration dominated these SR responses, increasing within hours, whereas root respiration responded more slowly over days. We conclude that root and microbial respiration sources respond differently in timing and magnitude to both seasonal and episodic moisture inputs. These findings have important implications for the mechanistic representation of SR in models and the response of dry ecosystems to changes in precipitation patterns.     

Light use efficiency of California redwood forest understory plants along a moisture gradient

We investigated photosynthesis of five plant species growing in the understory at three sites (1,170-, 1,600- and 2,100-mm annual moisture inputs), along the geographical range of coastal California redwood forest, to determine whether greater inputs of rain and fog at northern sites enhance photosynthetic utilization of fluctuating light. Measurements of understory light environment and gas exchange were carried out to determine steady state and dynamic photosynthetic responses to light. Leaf area index ranged from 4.84 at the 2,100-mm site to 5.98 at the 1,170-mm site. Maximum rates of net photosynthesis and stomatal conductance (g) did not vary appreciably within species across sites. Photosynthetic induction after a change from low to high light was significantly greater in plants growing in lower light conditions regardless of site. Photosynthetic induction also increased with the rate of g in diffuse light, prior to the increase to saturating light levels. Post-illumination CO₂ assimilation was the largest factor contributing to variation in C gain during simulated lightflecks. The duration of post-illumination photosynthetic activity, total CO₂ assimilation per light received, and light use efficiency during simulated lightflecks increased significantly with moisture inputs in four out of five species. Increasing leaf N concentration with increasing moisture inputs in three out of five species, coupled with changes in leaf N isotopic composition with the onset of the summer fog season suggest that natural N deposition increases with rain and fog inputs and contributes to greater utilization of fluctuating light availability in coastal California redwood forests.     

A decade of irrigation transforms the soil microbiome of a semi‐arid pine forest

The impact of climate change on the soil microbiome potentially alters the biogeochemical cycle of terrestrial ecosystems. In semi‐arid environments, water availability is a major constraint on biogeochemical cycles due to the combination of high summer temperatures and low rainfall. Here, we explored how 10 years of irrigation of a water‐limited pine forest in the central European Alps altered the soil microbiome and associated ecosystem functioning. A decade of irrigation stimulated tree growth, resulting in higher crown cover, larger yearly increments of tree biomass, increased litter fall and greater root biomass. Greater amounts of plant‐derived inputs associated with increased primary production in the irrigated forest stands stimulated soil microbial activity coupled with pronounced shifts in the microbiome from largely oligotrophic to more copiotrophic lifestyles. Microbial groups benefitting from increased resource availabilities (litter, rhizodeposits) thrived under irrigation, leading to enhanced soil organic matter mineralization and carbon respired from irrigated soils. This unique long‐term study provides new insights into the impact of precipitation changes on the soil microbiome and associated ecosystem functioning in a water‐limited pine forest ecosystem and improves our understanding of the persistency of long‐term soil carbon stocks in a changing climate.     

Fog Water and Ecosystem Function: Heterogeneity in a California Redwood Forest

Fog is thought to influence ecological function in coastal forests worldwide, yet few data are available that illuminate the mechanisms underlying this influence. In a California redwood forest we measured water and nitrogen (N) fluxes from horizontally moving fog and vertically delivered rain as well as redwood tree function. The spatial heterogeneity of water and N fluxes, water availability, tree water use, and canopy N processing varied greatly across seasons. Water and N fluxes to soil (annual average of 98% and 89%, respectively) across the whole forest occurred primarily in the rain season and was relatively even across the whole forest. In contrast, below-canopy flux of fog water and N declined exponentially from the windward edge to the forest interior. Following large fog events, soil moisture was greater at the windward edge than anywhere else in the forest. Physiological activity in redwoods reflected these differences in inputs across seasons: tree physiological responses did not vary spatially in the rain season, but in the fog season, water use was greater, yet water stress was less, in trees at the windward edge of the forest versus the interior. In both seasons, vertical passage through the forest changed the amount of water and form and concentration of N, revealing the role of the tree canopy in processing atmospheric inputs. Although total fog water inputs were comparatively small, they may have important ecosystem functions, including relief of canopy water stress and, where there is fog drip, functional coupling of above- and belowground processes. Author Contributions  We make no distinction in effort and contribution between the first and second authors or between the third and fourth authors. HAE and KCW were responsible for project design, oversight, data analysis, synthesis, and writing. PHT analyzed N samples and contributed significantly to interpretation of data. TED established the initial tree physiology study site, provided and analyzed the data on tree physiology, and contributed to the synthesis of these data. MKF first identified the lack of connection between inputs and microbial processing and catalyzed synthesis regarding belowground processing. AME created figures, performed field and laboratory work, and was database manager. VKSB collected field samples and carried out the litterfall study. All authors edited the paper.     

Maritime climate influence on chaparral composition and diversity in the coast range of central California

We investigated the hypothesis that maritime climatic factors associated with summer fog and low cloud stratus (summer marine layer) help explain the compositional diversity of chaparral in the coast range of central California. We randomly sampled chaparral species composition in 0.1‐hectare plots along a coast‐to‐interior gradient. For each plot, climatic variables were estimated and soil samples were analyzed. We used Cluster Analysis and Principle Components Analysis to objectively categorize plots into climate zone groups. Climate variables, vegetation composition and various diversity measures were compared across climate zone groups using ANOVA and nonmetric multidimensional scaling. Differences in climatic variables that relate to summer moisture availability and winter freeze events explained the majority of variance in measured conditions and coincided with three chaparral assemblages: maritime (lowland coast where the summer marine layer was strongest), transition (upland coast with mild summer marine layer influence and greater winter precipitation), and interior sites that generally lacked late summer water availability from either source. Species turnover (β‐diversity) was higher among maritime and transition sites than interior sites. Coastal chaparral differs from interior chaparral in having a higher obligate seeder to facultative seeder (resprouter) ratio and by being dominated by various Arctostaphylos species as opposed to the interior dominant, Adenostoma fasciculatum. The maritime climate influence along the California central coast is associated with patterns of woody plant composition and β‐diversity among sites. Summer fog in coastal lowlands and higher winter precipitation in coastal uplands combine to lower late dry season water deficit in coastal chaparral and contribute to longer fire return intervals that are associated with obligate seeders and more local endemism. Soil nutrients are comparatively less important in explaining plant community composition, but heterogeneous azonal soils contribute to local endemism and promote isolated chaparral patches within the dominant forest vegetation along the coast.     

Polystichum munitum (Dryopteridaceae) varies geographically in its capacity to absorb fog water by foliar uptake within the redwood forest ecosystem

? Premise of the study: Fog provides a critical water resource to plants around the world. In the redwood forest ecosystem of northern California, plants depend on fog absorbed through foliar uptake to stay hydrated during the rainless summer. In this study, we identified regions within the redwood ecosystem where the fern Polystichum munitum canopy most effectively absorbs fog drip that reaches the forest floor. ? Methods: We measured the foliar uptake capacity of P. munitum fronds at seven sites along 700 km of the redwood forest ecosystem. We quantified the canopy cover of P. munitum at each site and estimated how much water the fern canopy can acquire aboveground through fog interception and absorption. ? Key results: Throughout the ecosystem, nocturnal foliar uptake increased the leaf water content of P. munitum by 7.2%, and we estimated that the P. munitum canopy can absorb 5 ± 3% (mean ± SE) of intercepted fog precipitation. Strikingly, P. munitum had the highest foliar uptake capacity in the center of the ecosystem and may absorb 10% more of the fog its canopy intercepts in this region relative to other regions studied. Conversely, P. munitum had no foliar uptake capacity in the southern end of the ecosystem. ? Conclusions: This study shows the first evidence that foliar uptake varies within species at the landscape scale. Our findings suggest that the P. munitum at the southern tip of the redwood ecosystem may suffer most from low summertime water availability because it had no potential to acquire fog as an aboveground water subsidy.     

Ecohydrology of Adjacent Sagebrush and Lodgepole Pine Ecosystems: The Consequences of Climate Change and Disturbance

Sagebrush steppe and lodgepole pine forests are two of the most widespread vegetation types in the western United States and they play crucial roles in the hydrologic cycle of these water-limited regions. We used a process-based ecosystem water model to characterize the potential impact of climate change and disturbance (wildfire and beetle mortality) on water cycling in adjacent sagebrush and lodgepole pine ecosystems. Despite similar climatic and topographic conditions between these ecosystems at the sites examined, lodgepole pine, and sagebrush exhibited consistent differences in water balance, notably more evaporation and drier summer soils in the sagebrush and greater transpiration and less water yield in lodgepole pine. Canopy disturbances (either fire or beetle) have dramatic impacts on water balance and availability: reducing transpiration while increasing evaporation and water yield. Results suggest that climate change may reduce snowpack, increase evaporation and transpiration, and lengthen the duration of dry soil conditions in the summer, but may have uncertain effects on drainage. Changes in the distribution of sagebrush and lodgepole pine ecosystems as a consequence of climate change and/or altered disturbance regimes will likely alter ecosystem water balance.     

Soil Microbial Community Response to Drought and Precipitation Variability in the Chihuahuan Desert

Increases in the magnitude and variability of precipitation events have been predicted for the Chihuahuan Desert region of West Texas. As patterns of moisture inputs and amounts change, soil microbial communities will respond to these alterations in soil moisture windows. In this study, we examined the soil microbial community structure within three vegetation zones along the Pine Canyon Watershed, an elevation and vegetation gradient in Big Bend National Park, Chihuahuan Desert. Soil samples at each site were obtained in mid-winter (January) and in mid-summer (August) for 2 years to capture a component of the variability in soil temperature and moisture that can occur seasonally and between years along this watershed. Precipitation patterns and amounts differed substantially between years with a drought characterizing most of the second year. Soils were collected during the drought period and following a large rainfall event and compared to soil samples collected during a relatively average season. Structural changes within microbial community in response to site, season, and precipitation patterns were evaluated using fatty acid methyl ester (FAME) and polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE) analyses. Fungal FAME amounts differed significantly across seasons and sites and greatly outweighed the quantity of bacterial and actinomycete FAME levels for all sites and seasons. The highest fungal FAME levels were obtained in the low desert scrub site and not from the high elevation oak–pine forests. Total bacterial and actinomycete FAME levels did not differ significantly across season and year within any of the three locations along the watershed. Total bacterial and actinomycete FAME levels in the low elevation desert-shrub and grassland sites were slightly higher in the winter than in the summer. Microbial community structure at the high elevation oak–pine forest site was strongly correlated with levels of NH₄ ⁺–N, % soil moisture, and amounts of soil organic matter irrespective of season. Microbial community structure at the low elevation desert scrub and sotol grasslands sites was most strongly related to soil pH with bacterial and actinobacterial FAME levels accounting for site differences along the gradient. DGGE band counts of amplified soil bacterial DNA were found to differ significantly across sites and season with the highest band counts found in the mid-elevation grassland site. The least number of bands was observed in the high elevation oak–pine forest following the large summer-rain event that occurred after a prolonged drought. Microbial responses to changes in precipitation frequency and amount due to climate change will differ among vegetation zones along this Chihuahuan Desert watershed gradient. Soil bacterial communities at the mid-elevation grasslands site are the most vulnerable to changes in precipitation frequency and timing, while fungal community structure is most vulnerable in the low desert scrub site. The differential susceptibility of the microbial communities to changes in precipitation amounts along the elevation gradient reflects the interactive effects of the soil moisture window duration following a precipitation event and differences in soil heat loads. Amounts and types of carbon inputs may not be as important in regulating microbial structure among vegetation zones within in an arid environment as is the seasonal pattern of soil moisture and the soil heat load profile that characterizes the location.     

Salt marsh as a coastal filter for the oceans: changes in function with experimental increases in nitrogen loading and sea-level rise

Coastal salt marshes are among Earth's most productive ecosystems and provide a number of ecosystem services, including interception of watershed-derived nitrogen (N) before it reaches nearshore oceans. Nitrogen pollution and climate change are two dominant drivers of global-change impacts on ecosystems, yet their interacting effects at the land-sea interface are poorly understood. We addressed how sea-level rise and anthropogenic N additions affect the salt marsh ecosystem process of nitrogen uptake using a field-based, manipulative experiment. We crossed simulated sea-level change and ammonium-nitrate (NH(4)NO(3))-addition treatments in a fully factorial design to examine their potentially interacting effects on emergent marsh plants in a central California estuary. We measured above- and belowground biomass and tissue nutrient concentrations seasonally and found that N-addition had a significant, positive effect on a) aboveground biomass, b) plant tissue N concentrations, c) N stock sequestered in plants, and d) shoot:root ratios in summer. Relative sea-level rise did not significantly affect biomass, with the exception of the most extreme sea-level-rise simulation, in which all plants died by the summer of the second year. Although there was a strong response to N-addition treatments, salt marsh responses varied by season. Our results suggest that in our site at Coyote Marsh, Elkhorn Slough, coastal salt marsh plants serve as a robust N trap and coastal filter; this function is not saturated by high background annual N inputs from upstream agriculture. However, if the marsh is drowned by rising seas, as in our most extreme sea-level rise treatment, marsh plants will no longer provide the ecosystem service of buffering the coastal ocean from eutrophication.     

Medium,Vector, and Connector: Fog and the Maintenance of Ecosystems

Ecosystems - Fog and low-lying cloud (fog) play a significant role in the maintenance of ecosystems, from desert to alpine and from coastal to inland systems. Our central thesis is that fog...     

Do soil enzymes respond to long-term grazing in an arid ecosystem?

Background and aims

Our objective was to assess the effects of long-term continuous grazing on soil enzyme activities in relation to shifts in plant litter attributes and soil resources in an arid ecosystem, considering both spatial and temporal variations.

Methods

We randomly extracted soil samples with the respective litter cover at 5 modal size plant-covered patches (PCP) and the nearest inter-canopy areas (IC) at Patagonian Monte sites with low, medium and high grazing intensity in winter and summer from 2007 to 2009. We analyzed enzyme activities (dehydrogenase, ß-glucosidase, protease, alkaline and acid phosphatase), microbial biomass-C, organic-C, total soil-N, and moisture in soil and mass and quality in plant litter. We assessed faeces density and plant cover in the field.

Results and conclusions

Grazing led to reduced grass cover, decreasing plant litter mass with increasing soluble phenolics, and reduced phosphatases, ß-glucosidase and microbial biomass-C at PCP. A localized nutrient input from animal excreta seems to promote microbial biomass-C, alkaline phosphatase and dehydrogenase activities but only at IC from the site with high grazing intensity. Plant heterogeneous distribution, plant litter quantity and quality, nutrient inputs from grazers and seasonal variation in soil moisture, also affecting soil resources and microbial biomass, modulate soil enzyme responses to long-term grazing in the arid Patagonian Monte.     

Laserfarm – A high-throughput workflow for generating geospatial data products of ecosystem structure from airborne laser scanning point clouds

Quantifying ecosystem structure is of key importance for ecology, conservation, restoration, and biodiversity monitoring because the diversity, geographic distribution and abundance of animals, plants and other organisms is tightly linked to the physical structure of vegetation and associated microclimates. Light Detection And Ranging (LiDAR) — an active remote sensing technique — can provide detailed and high resolution information on ecosystem structure because the laser pulse emitted from the sensor and its subsequent return signal from the vegetation (leaves, branches, stems) delivers three-dimensional point clouds from which metrics of vegetation structure (e.g. ecosystem height, cover, and structural complexity) can be derived. However, processing 3D LiDAR point clouds into geospatial data products of ecosystem structure remains challenging across broad spatial extents due to the large volume of national or regional point cloud datasets (typically multiple terabytes consisting of hundreds of billions of points). Here, we present a high-throughput workflow called ‘Laserfarm’ enabling the efficient, scalable and distributed processing of multi-terabyte LiDAR point clouds from national and regional airborne laser scanning (ALS) surveys into geospatial data products of ecosystem structure. Laserfarm is a free and open-source, end-to-end workflow which contains modular pipelines for the re-tiling, normalization, feature extraction and rasterization of point cloud information from ALS and other LiDAR surveys. The workflow is designed with horizontal scalability and can be deployed with distributed computing on different infrastructures, e.g. a cluster of virtual machines. We demonstrate the Laserfarm workflow by processing a country-wide multi-terabyte ALS dataset of the Netherlands (covering ∼34,000 km² with ∼700 billion points and ∼ 16 TB uncompressed LiDAR point clouds) into 25 raster layers at 10 m resolution capturing ecosystem height, cover and structural complexity at a national extent. The Laserfarm workflow, implemented in Python and available as Jupyter Notebooks, is applicable to other LiDAR datasets and enables users to execute automated pipelines for generating consistent and reproducible geospatial data products of ecosystems structure from massive amounts of LiDAR point clouds on distributed computing infrastructures, including cloud computing environments. We provide information on workflow performance (including total CPU times, total wall-time estimates and average CPU times for single files and LiDAR metrics) and discuss how the Laserfarm workflow can be scaled to other LiDAR datasets and computing environments, including remote cloud infrastructures. The Laserfarm workflow allows a broad user community to process massive amounts of LiDAR point clouds for mapping vegetation structure, e.g. for applications in ecology, biodiversity monitoring and ecosystem restoration.     

Season mediates herbivore effects on litter and soil microbial abundance and activity in a semi-arid woodland

Herbivores can directly impact ecosystem function by altering litter quality of an ecosystem or indirectly by shifting the composition of microbial communities that mediate nutrient processes. We examined the effects of tree susceptibility and resistance to herbivory on litter microarthropod and soil microbial communities to test the general hypothesis that herbivore driven changes in litter inputs and soil microclimate will feedback to the microbial community. Our study population consisted of individual piñon pine trees that were either susceptible or resistant to the stem-boring moth (Dioryctria albovittella) and susceptible piñon pine trees from which the moth herbivores have been manually removed since 1982. Moth herbivory increased piñon litter nitrogen concentrations (16%) and decreased canopy precipitation interception (28%), both potentially significant factors influencing litter and soil microbial communities. Our research resulted in three major findings: (1) In spite of an apparent increase in litter quality, herbivory did not change litter microarthropod abundance or species richness. (2) However, susceptibility to herbivores strongly influenced bulk soil microbial communities (i.e., 52% greater abundance beneath herbivore-resistant and herbivore-removal trees than susceptible trees) and alkaline phosphatase activity (i.e., 412% increase beneath susceptible trees relative to other groups). (3) Season had a strong influence on microbial communities (i.e., microbial biomass and alkaline phosphatase activity increased after the summer rains), and their response to herbivore inputs, in this semi-arid ecosystem. Thus, during the dry season plant resistance and susceptibility to a common insect herbivore had little or no observable effects on the belowground organisms and processes we studied, but after the rains, some pronounced effects emerged.     

The contribution of fog to the water relations of Sequoia sempervirens (D. Don): foliar uptake and prevention of dehydration

Fog is a defining feature of the coastal California redwood forest and fog inputs via canopy drip in summer can constitute 30% or more of the total water input each year. A great deal of occult precipitation (fog and light rain) is retained in redwood canopies, which have some of the largest leaf area indices known (Westman & Whittaker, Journal of Ecology 63, 493–520, 1975). An investigation was carried out to determine whether some fraction of intercepted fog water might be directly absorbed through leaf surfaces and if so, the importance of this to the water relations physiology of coast redwood, Sequoia sempervirens. An array of complimentary techniques were adopted to demonstrate that fog is absorbed directly by S. sempervirens foliage. Xylem sap transport reversed direction during heavy fog, with instantaneous flow rates in the direction of the soil peaking at approximately 5–7% of maximum transpiration rate. Isotopic analyses showed that up to 6% of a leaf's water content could be traced to a previous night's fog deposition, but this amount varied considerably depending on the age and water status of the leaves. Old leaves, which appear most able to absorb fog water were able to absorb distilled water when fully submersed at an average rate of 0.90 mmol m² s^?1, or about 80% of transpiration rates measured at the leaf level in the field. Sequoia sempervirens has poor stomatal control in response to a drying atmosphere, with rates of water loss on very dry nights up to 40% of midday summer values and rates above 10% being extremely common. Owing to this profligate water use behaviour of S. sempervirens, it appears that fog has a greater role in suppressing water loss from leaves, and thereby ameliorating daily water stress, than in providing supplemental water to foliar tissues per se. Although direct foliar absorption from fog inputs represents only a small fraction of the water used each day, fog's in reducing transpiration and rehydrating leaf tissues during the most active growth periods in summer may allow for greater seasonal carbon fixation and thus contribute to the very fast growth rates and great size of this species.     

Summer water use by California coastal prairie grasses: fog,drought, and community composition

Plants in the Mediterranean climate region of California typically experience summer drought conditions, but correlations between zones of frequent coastal fog inundation and certain species’ distributions suggest that water inputs from fog may influence species composition in coastal habitats. We sampled the stable H and O isotope ratios of water in non-photosynthetic plant tissue from a variety of perennial grass species and soil in four sites in northern California in order to determine the proportion of water deriving from winter rains and fog during the summer. The relationship between H and O stable isotopes from our sample sites fell to the right of the local meteoric water line (LMWL) during the summer drought, providing evidence that evaporation of water from the soil had taken place prior to the uptake of water by vegetation. We developed a novel method to infer the isotope values of water before it was subjected to evaporation in which we used experimental data to calculate the slope of the δH versus δO line versus the LMWL. After accounting for evaporation, we then used a two-source mixing model to evaluate plant usage of fog water. The model indicated that 28–66% of the water taken up by plants via roots during the summer drought came from fog rather than residual soil water from winter rain. Fog use decreased as distance from the coast increased, and there were significant differences among species in the use of fog. Rather than consistent differences in fog use by species whose distributions are limited to the coast versus those with broader distributions, species responded individualistically to summer fog. We conclude that fogwater inputs can mitigate the summer drought in coastal California for many species, likely giving an advantage to species that can use it over species that cannot.     

Soil animal responses to moisture availability are largely scale,not ecosystem dependent: insight from a cross‐site study

Climate change will result in reduced soil water availability in much of the world either due to changes in precipitation or increased temperature and evapotranspiration. How communities of mites and nematodes may respond to changes in moisture availability is not well known, yet these organisms play important roles in decomposition and nutrient cycling processes. We determined how communities of these organisms respond to changes in moisture availability and whether common patterns occur along fine‐scale gradients of soil moisture within four individual ecosystem types (mesic, xeric and arid grasslands and a polar desert) located in the western United States and Antarctica, as well as across a cross‐ecosystem moisture gradient (CEMG) of all four ecosystems considered together. An elevation transect of three sampling plots was monitored within each ecosystem and soil samples were collected from these plots and from existing experimental precipitation manipulations within each ecosystem once in fall of 2009 and three times each in 2010 and 2011. Mites and nematodes were sorted to trophic groups and analyzed to determine community responses to changes in soil moisture availability. We found that while both mites and nematodes increased with available soil moisture across the CEMG, within individual ecosystems, increases in soil moisture resulted in decreases to nematode communities at all but the arid grassland ecosystem; mites showed no responses at any ecosystem. In addition, we found changes in proportional abundances of mite and nematode trophic groups as soil moisture increased within individual ecosystems, which may result in shifts within soil food webs with important consequences for ecosystem functioning. We suggest that communities of soil animals at local scales may respond predictably to changes in moisture availability regardless of ecosystem type but that additional factors, such as climate variability, vegetation composition, and soil properties may influence this relationship over larger scales.     

Light‐driven tipping points in polar ecosystems

Some ecosystems can undergo abrupt transformation in response to relatively small environmental change. Identifying imminent ‘tipping points’ is crucial for biodiversity conservation, particularly in the face of climate change. Here, we describe a tipping point mechanism likely to induce widespread regime shifts in polar ecosystems. Seasonal snow and ice‐cover periodically block sunlight reaching polar ecosystems, but the effect of this on annual light depends critically on the timing of cover within the annual solar cycle. At high latitudes, sunlight is strongly seasonal, and ice‐free days around the summer solstice receive orders of magnitude more light than those in winter. Early melt that brings the date of ice‐loss closer to midsummer will cause an exponential increase in the amount of sunlight reaching some ecosystems per year. This is likely to drive ecological tipping points in which primary producers (plants and algae) flourish and out‐compete dark‐adapted communities. We demonstrate this principle on Antarctic shallow seabed ecosystems, which our data suggest are sensitive to small changes in the timing of sea‐ice loss. Algae respond to light thresholds that are easily exceeded by a slight reduction in sea‐ice duration. Earlier sea‐ice loss is likely to cause extensive regime shifts in which endemic shallow‐water invertebrate communities are replaced by algae, reducing coastal biodiversity and fundamentally changing ecosystem functioning. Modeling shows that recent changes in ice and snow cover have already transformed annual light budgets in large areas of the Arctic and Antarctic, and both aquatic and terrestrial ecosystems are likely to experience further significant change in light. The interaction between ice‐loss and solar irradiance renders polar ecosystems acutely vulnerable to abrupt ecosystem change, as light‐driven tipping points are readily breached by relatively slight shifts in the timing of snow and ice‐loss.